新疆农业科学 ›› 2024, Vol. 61 ›› Issue (2): 421-433.DOI: 10.6048/j.issn.1001-4330.2024.02.018
孟涵颖(), 郭丹丹, 何婉洁, 张伟伟, 张建萍, 陈静(
)
收稿日期:
2023-06-02
出版日期:
2024-02-20
发布日期:
2024-03-19
通信作者:
陈静(1979-),女,陕西人,教授,博士,硕士生导师,研究方向为农业昆虫与害虫防治,(E-mail)Chj_agr@shzu.edu.cn作者简介:
孟涵颖(1997-),女,新疆塔城人,硕士研究生,研究方向为农业昆虫与害虫防治,(E-mail)1174561906@qq.com
基金资助:
MENG Hanying(), GUO Dandan, HE Wanjie, ZHANG Weiwei, ZHANG Jianping, CHEN Jing(
)
Received:
2023-06-02
Published:
2024-02-20
Online:
2024-03-19
Supported by:
摘要:
【目的】分析双斑长跗萤叶甲取食前后的玉米叶片挥发物组成和含量,为进一步研究玉米叶片挥发物对双斑萤叶甲的防御机制提供科学依据。【方法】采用固相微萃取和气相色谱-质谱联用仪检测并分析健康玉米叶片及双斑长跗萤叶甲不同取食时间(6、24、36和48 h)下玉米叶片挥发物组分变化。【结果】双斑长跗萤叶甲取食0、6、24、36和48 h后,分别检测到29、30、33、36、26种挥发物,主要为醛类、萜类、醇类及酮类等化合物。1-石竹烯、顺-11-十六醛、α-石竹烯、正己醛等20种挥发物随该虫取食为害后消失;新收集到2-己烯醛、(E)-2-己烯醛、α-蒎烯、β-月桂烯等42种化合物。α-蒎烯、2-己烯醛、β-蒎烯等物质是虫害6及24 h释放的主要化合物;十六醛、十四醛、正十五碳醛、β-紫罗酮、(E)-3-己烯-1-醇是虫害36 h释放的主要化合物;β-红没药醇、橙花叔醇、(1R)-α-蒎烯、β-罗勒烯、β-檀香烯、β-月桂烯、葎草烯是虫害48 h释放的主要化合物。【结论】双斑长跗萤叶甲不同时间取食玉米叶片的挥发物的组分和含量影响较大,取食6及24 h主要释放烯烃类化合物,取食36 h主要释放醛类和酮类化合物,取食48 h主要释放醇类、烯烃类化合物。
中图分类号:
孟涵颖, 郭丹丹, 何婉洁, 张伟伟, 张建萍, 陈静. 双斑长跗萤叶甲取食玉米叶片的挥发物分析[J]. 新疆农业科学, 2024, 61(2): 421-433.
MENG Hanying, GUO Dandan, HE Wanjie, ZHANG Weiwei, ZHANG Jianping, CHEN Jing. Effects of different feeding time of Monolepta Signata on the composition and content of volatile in maize leaves[J]. Xinjiang Agricultural Sciences, 2024, 61(2): 421-433.
图1 罩笼示意(左)和被双斑长跗萤叶甲分别取食6、24、36和48 h后的玉米叶片(右)
Fig.1 Schematic diagram of the cage(left)and maize leaves after 6, 24, 36 and 48 h of feeding induction by M.signata(right)
保留时间 Retention time (min) | 物质名称 Substance name | CAS号 CAS Number | 相对含量 Relative content (%) |
---|---|---|---|
3.68 | 正己醛 | 66-25-1 | 0.17 |
3.74 | 异丁烯 | 115-11-7 | 0.5 |
4.78 | 1,3-丁二烯醇 | 106-99-0 | 0.38 |
4.83 | 2,3-二甲基- 1,3-丁二烯 | 151832-72-3 | 0.43 |
7.26 | 3-已基1,5-辛二烯 | 111-78-4 | 0.55 |
8.99 | 1,6-辛二烯 | 3710-41-6 | 0.7 |
16.93 | 1,3,4-三甲基- 3环己烯-1-甲醛 | 40702-26-9 | 0.38 |
22.69 | 5-十二烯 | 7206-28-2 | 0.41 |
22.94 | 正十四烷 | 629-59-4 | 0.44 |
23.59 | 1-石竹烯 | 87-44-5 | 12.49 |
23.85 | 氧化石竹烯 | 1139-30-6 | 0.76 |
24.64 | α-石竹烯 | 6753-98-6 | 0.85 |
25.63 | β-紫罗酮 | 79-77-6 | 3.81 |
26.03 | 正十五烷 | 629-62-9 | 0.43 |
26.35 | β-红没药醇 | 495-61-4 | 0.37 |
28.52 | 反-Zα环氧化红没药烯 | 515-69-5 | 1.73 |
28.96 | 正十六烷 | 544-76-3 | 0.37 |
29.31 | 7-十三烯醇(Z) | 74962-99-5 | 2.05 |
31.21 | (Z)-9-肉豆蔻醛 | 124-25-4 | 0.21 |
31.32 | 正十七烷 | 629-78-7 | 0.2 |
31.64 | 正十五碳醛 | 2765-11-9 | 66.95 |
32.64 | (Z)-9-十六碳烯醛 | 56219-04-6 | 0.36 |
33.29 | 环庚醇 | 502-41-0 | 0.3 |
33.6 | 植物醇 | 150-86-7 | 1.31 |
34.03 | 邻苯二甲酸二异丁酯 | 84-69-5 | 1.97 |
34.36 | 顺-11-十六醛 | 53939-28-9 | 0.17 |
35.17 | 棕榈酸 | 57-10-3 | 0.63 |
37.64 | 2,6,6-三甲基3-(2-丙 烯基)双环[3.1.1]庚烷 | 4889-83-2 | 0.79 |
37.68 | 2,5-二甲基四氢- 2-甲醇吡喃 | 54004-46-5 | 0.29 |
表1 玉米健康叶片挥发物的化学成分
Tab.1 Chemical composition of volatiles from healthy maize leaves
保留时间 Retention time (min) | 物质名称 Substance name | CAS号 CAS Number | 相对含量 Relative content (%) |
---|---|---|---|
3.68 | 正己醛 | 66-25-1 | 0.17 |
3.74 | 异丁烯 | 115-11-7 | 0.5 |
4.78 | 1,3-丁二烯醇 | 106-99-0 | 0.38 |
4.83 | 2,3-二甲基- 1,3-丁二烯 | 151832-72-3 | 0.43 |
7.26 | 3-已基1,5-辛二烯 | 111-78-4 | 0.55 |
8.99 | 1,6-辛二烯 | 3710-41-6 | 0.7 |
16.93 | 1,3,4-三甲基- 3环己烯-1-甲醛 | 40702-26-9 | 0.38 |
22.69 | 5-十二烯 | 7206-28-2 | 0.41 |
22.94 | 正十四烷 | 629-59-4 | 0.44 |
23.59 | 1-石竹烯 | 87-44-5 | 12.49 |
23.85 | 氧化石竹烯 | 1139-30-6 | 0.76 |
24.64 | α-石竹烯 | 6753-98-6 | 0.85 |
25.63 | β-紫罗酮 | 79-77-6 | 3.81 |
26.03 | 正十五烷 | 629-62-9 | 0.43 |
26.35 | β-红没药醇 | 495-61-4 | 0.37 |
28.52 | 反-Zα环氧化红没药烯 | 515-69-5 | 1.73 |
28.96 | 正十六烷 | 544-76-3 | 0.37 |
29.31 | 7-十三烯醇(Z) | 74962-99-5 | 2.05 |
31.21 | (Z)-9-肉豆蔻醛 | 124-25-4 | 0.21 |
31.32 | 正十七烷 | 629-78-7 | 0.2 |
31.64 | 正十五碳醛 | 2765-11-9 | 66.95 |
32.64 | (Z)-9-十六碳烯醛 | 56219-04-6 | 0.36 |
33.29 | 环庚醇 | 502-41-0 | 0.3 |
33.6 | 植物醇 | 150-86-7 | 1.31 |
34.03 | 邻苯二甲酸二异丁酯 | 84-69-5 | 1.97 |
34.36 | 顺-11-十六醛 | 53939-28-9 | 0.17 |
35.17 | 棕榈酸 | 57-10-3 | 0.63 |
37.64 | 2,6,6-三甲基3-(2-丙 烯基)双环[3.1.1]庚烷 | 4889-83-2 | 0.79 |
37.68 | 2,5-二甲基四氢- 2-甲醇吡喃 | 54004-46-5 | 0.29 |
化合物 种类 Compound type | 化合物名称 Compound name | 序号 Number | CAS号 CAS Number | 相对含量 Relative content(%) | |||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
CK | 6 h | 24 h | 36 h | 48 h | |||||||||||||
醇类 Alcohols | 3,7,11-三甲基-1-十二烷醇 | v1 | 6750-34-1 | - | - | - | 1.09 | - | |||||||||
(E)-2-十四碳烯-1-醇 | v2 | 35153-18-5 | - | - | - | 0.55 | - | ||||||||||
3,7,7三甲基双环[2.2.1] 庚环-2-甲基醇 | v3 | 128376-90-9 | - | - | - | 0.76 | - | ||||||||||
D-橙花叔醇 | v4 | 3790-78-1 | - | 0.52 | 0.15 | - | 0.09 | ||||||||||
β-红没药醇 | v5 | 495-61-4 | 0.37 | 9.17 | 7.44 | - | 7.71 | ||||||||||
6,11-二甲基-2,6, 10-十二碳三烯-1-醇 | v6 | 4602-84-0 | - | - | - | 1.85 | - | ||||||||||
δ-杜松醇 | v7 | 17910-08-6 | - | - | - | 0.96 | - | ||||||||||
橙花叔醇 | v8 | 7212-44-4 | - | - | 0.28 | 0 | 0.19 | ||||||||||
植醇 | v9 | 7541-49-3 | 1.31 | - | - | 0.59 | - | ||||||||||
异植物醇 | v10 | 505-32-8 | - | - | - | 0.68 | - | ||||||||||
萜类 Terpenoids | (1R)-(+)-α蒎烯 | v11 | 70750-57-1 | - | 4.85 | 2.28 | - | - | |||||||||
2,3-二甲基-2-降冰片烯 | v12 | 694-92-8 | - | - | - | - | 0.08 | ||||||||||
(E)-β-法呢烯 | v13 | 18794-84-8 | - | - | - | - | 0.36 | ||||||||||
D-柠檬烯 | v14 | 5989-27-5 | - | 0.19 | 0.15 | - | 0.19 | ||||||||||
L-柠檬烯 | v15 | 8050-32-6 | - | - | - | - | 0.08 | ||||||||||
α-荜澄茄油烯 | v16 | 17699-14-8 | - | 0.99 | 0.81 | - | - | ||||||||||
α-蒎烯 | v17 | 7785-26-4 | - | - | 10.21 | 2.59 | 0.74 | ||||||||||
β-杜松 | v18 | 483-76-1 | - | 1.13 | 0.92 | - | 0.15 | ||||||||||
β-罗勒烯 | v19 | 13877-91-3 | - | - | - | - | 0.36 | ||||||||||
β-蒎烯 | v20 | 127-91-3 | - | 0.82 | 2.43 | - | 0.89 | ||||||||||
顺γ-红没药烯 | v28 | 495-62-5 | - | 16.61 | 13.47 | 1.29 | 20.14 | ||||||||||
新植二烯 | v29 | 504-96-1 | - | - | - | 1.36 | - | ||||||||||
氧化石竹烯 | v30 | 1139-30-6 | 0.76 | 0.52 | 0.42 | - | - | ||||||||||
榄香烯异构体 | v31 | 502-61-4 | - | 1.75 | 1.42 | - | 0.23 | ||||||||||
葎草烯 | v32 | 6753-98-6 | - | 14.46 | 11.72 | 1.76 | 12.83 | ||||||||||
绿叶化合物 Green leaf compounds | 3,5,5-三甲基-环己烯 | v33 | 933-120 | - | 0.1 | - | - | - | |||||||||
1,5,5-三甲基- 6-亚甲基环己烯 | v34 | 16609-28-2 | - | 2.69 | 2.18 | - | 0.59 | ||||||||||
2-己烯醛 | v35 | 6728-26-3 | - | 0.79 | 6.02 | - | 0.86 | ||||||||||
(E)-3-己烯-1-醇 | v36 | 3681-82-1 | - | - | 1.05 | 1.28 | - | ||||||||||
(Z)-3-己烯-1-醇 | v37 | 928-96-1 | - | - | - | 1.41 | - | ||||||||||
丁烯基环己烷 | v38 | 18736-82-8 | - | 0.12 | 0.09 | 0 | - | ||||||||||
酯类 Esters | 邻苯二甲酸丁酯 | v39 | 84-74-2 | - | - | - | 0.83 | - | |||||||||
邻苯二甲酸二异丁酯 | v40 | 84-69-5 | 1.97 | - | - | 1.37 | - | ||||||||||
12-氧代-十八碳-9-甲酸甲酯 | v41 | 2380-27-0 | - | - | - | 0.35 | - | ||||||||||
2-甲基-壬基丙酸酯 | v42 | 65833-30-9 | - | 0.1 | 0.08 | 0 | - | ||||||||||
3-甲基甲基丙烯酸戊酯 | v43 | 2849-98-1 | - | 0 | 0.16 | 0 | - | ||||||||||
肌醇六乙酸酯 | v44 | 1254-38-2 | - | - | - | 2.57 | - | ||||||||||
甲基丙烯酸3-甲基戊酯 | v45 | 541-47-9 | - | 0.1 | - | 0 | - | ||||||||||
顺-9-十四碳烯酸异丁酯 | v46 | 16725-53-4 | - | 0 | - | 0.55 | - | ||||||||||
十二酸乙酯 | v47 | 18281-05-5 | - | 0.07 | 0.06 | 0 | - | ||||||||||
醛类 Aldehyde | 13-甲基十四醛 | v48 | 2485-71-4 | - | - | - | 1.09 | - | |||||||||
13-十八碳烯醛 | v49 | 58594-45-9 | - | - | - | 0.42 | - | ||||||||||
3-甲基-戊醛 | v50 | 15877-57-3 | - | - | - | 0 | 0.13 | ||||||||||
十六烷醛 | v51 | 629-80-1 | - | - | - | 0.57 | - | ||||||||||
十四醛 | v52 | 124-25-4 | - | - | - | 4.74 | - | ||||||||||
顺-13-十八烯醛 | v53 | 58594-45-9 | - | - | - | 0.71 | - | ||||||||||
正十五碳醛 | v54 | 2756-11-9 | 66.95 | 0.29 | 0.24 | 48.39 | 0.07 | ||||||||||
烷烃类 Alkanes | 十七烷 | v55 | 629-78-7 | - | - | - | 0.94 | - | |||||||||
2,6,10,15-四甲基-十七烷 | v56 | 18344-37-1 | - | - | - | 1.41 | - | ||||||||||
2-甲基-十九烷 | v57 | 1560-86-7 | - | - | - | 0.41 | - | ||||||||||
10,10-二甲基-2,6-二亚甲 二环[7.2.0]十一烷 | v58 | 87399-97-1 | - | 1.67 | 1.35 | - | 2.23 | ||||||||||
十四烷 | v59 | 629-59-4 | 0.44 | - | - | 1.62 | - | ||||||||||
十五烷 | v60 | 629-62-9 | 0.43 | - | - | 1.02 | - | ||||||||||
十六烷 | v61 | 544-76-3 | 0.37 | - | - | 2.15 | - | ||||||||||
酮类 Ketones | β-紫罗酮 | v62 | 79-77-6 | 3.81 | - | - | 3.02 | - | |||||||||
4-异丙基-5,10-二甲基 十氢萘-1,3-二酮 | v63 | 37830-90-3 | - | 0.1 | 0.09 | 0 | - | ||||||||||
5,6-环氧-β紫罗兰酮 | v64 | 20483-36-7 | - | - | - | 0.69 | - | ||||||||||
6,10,14-三甲基-2-十五烷酮 | v65 | 502-69-2 | - | - | - | 0.5 | - | ||||||||||
(Z)--香叶基丙酮 | v66 | 3796-70-1 | - | - | - | 0.65 | - | ||||||||||
酰胺类Amides | 己内酰胺 | v67 | 105-60-2 | - | - | - | 0.92 | - |
表2 双斑长跗萤叶甲取食后玉米叶片释放的化合物相对含量变化
Tab.2 Compound content released from maize leaves after feeding by the M.signata
化合物 种类 Compound type | 化合物名称 Compound name | 序号 Number | CAS号 CAS Number | 相对含量 Relative content(%) | |||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
CK | 6 h | 24 h | 36 h | 48 h | |||||||||||||
醇类 Alcohols | 3,7,11-三甲基-1-十二烷醇 | v1 | 6750-34-1 | - | - | - | 1.09 | - | |||||||||
(E)-2-十四碳烯-1-醇 | v2 | 35153-18-5 | - | - | - | 0.55 | - | ||||||||||
3,7,7三甲基双环[2.2.1] 庚环-2-甲基醇 | v3 | 128376-90-9 | - | - | - | 0.76 | - | ||||||||||
D-橙花叔醇 | v4 | 3790-78-1 | - | 0.52 | 0.15 | - | 0.09 | ||||||||||
β-红没药醇 | v5 | 495-61-4 | 0.37 | 9.17 | 7.44 | - | 7.71 | ||||||||||
6,11-二甲基-2,6, 10-十二碳三烯-1-醇 | v6 | 4602-84-0 | - | - | - | 1.85 | - | ||||||||||
δ-杜松醇 | v7 | 17910-08-6 | - | - | - | 0.96 | - | ||||||||||
橙花叔醇 | v8 | 7212-44-4 | - | - | 0.28 | 0 | 0.19 | ||||||||||
植醇 | v9 | 7541-49-3 | 1.31 | - | - | 0.59 | - | ||||||||||
异植物醇 | v10 | 505-32-8 | - | - | - | 0.68 | - | ||||||||||
萜类 Terpenoids | (1R)-(+)-α蒎烯 | v11 | 70750-57-1 | - | 4.85 | 2.28 | - | - | |||||||||
2,3-二甲基-2-降冰片烯 | v12 | 694-92-8 | - | - | - | - | 0.08 | ||||||||||
(E)-β-法呢烯 | v13 | 18794-84-8 | - | - | - | - | 0.36 | ||||||||||
D-柠檬烯 | v14 | 5989-27-5 | - | 0.19 | 0.15 | - | 0.19 | ||||||||||
L-柠檬烯 | v15 | 8050-32-6 | - | - | - | - | 0.08 | ||||||||||
α-荜澄茄油烯 | v16 | 17699-14-8 | - | 0.99 | 0.81 | - | - | ||||||||||
α-蒎烯 | v17 | 7785-26-4 | - | - | 10.21 | 2.59 | 0.74 | ||||||||||
β-杜松 | v18 | 483-76-1 | - | 1.13 | 0.92 | - | 0.15 | ||||||||||
β-罗勒烯 | v19 | 13877-91-3 | - | - | - | - | 0.36 | ||||||||||
β-蒎烯 | v20 | 127-91-3 | - | 0.82 | 2.43 | - | 0.89 | ||||||||||
顺γ-红没药烯 | v28 | 495-62-5 | - | 16.61 | 13.47 | 1.29 | 20.14 | ||||||||||
新植二烯 | v29 | 504-96-1 | - | - | - | 1.36 | - | ||||||||||
氧化石竹烯 | v30 | 1139-30-6 | 0.76 | 0.52 | 0.42 | - | - | ||||||||||
榄香烯异构体 | v31 | 502-61-4 | - | 1.75 | 1.42 | - | 0.23 | ||||||||||
葎草烯 | v32 | 6753-98-6 | - | 14.46 | 11.72 | 1.76 | 12.83 | ||||||||||
绿叶化合物 Green leaf compounds | 3,5,5-三甲基-环己烯 | v33 | 933-120 | - | 0.1 | - | - | - | |||||||||
1,5,5-三甲基- 6-亚甲基环己烯 | v34 | 16609-28-2 | - | 2.69 | 2.18 | - | 0.59 | ||||||||||
2-己烯醛 | v35 | 6728-26-3 | - | 0.79 | 6.02 | - | 0.86 | ||||||||||
(E)-3-己烯-1-醇 | v36 | 3681-82-1 | - | - | 1.05 | 1.28 | - | ||||||||||
(Z)-3-己烯-1-醇 | v37 | 928-96-1 | - | - | - | 1.41 | - | ||||||||||
丁烯基环己烷 | v38 | 18736-82-8 | - | 0.12 | 0.09 | 0 | - | ||||||||||
酯类 Esters | 邻苯二甲酸丁酯 | v39 | 84-74-2 | - | - | - | 0.83 | - | |||||||||
邻苯二甲酸二异丁酯 | v40 | 84-69-5 | 1.97 | - | - | 1.37 | - | ||||||||||
12-氧代-十八碳-9-甲酸甲酯 | v41 | 2380-27-0 | - | - | - | 0.35 | - | ||||||||||
2-甲基-壬基丙酸酯 | v42 | 65833-30-9 | - | 0.1 | 0.08 | 0 | - | ||||||||||
3-甲基甲基丙烯酸戊酯 | v43 | 2849-98-1 | - | 0 | 0.16 | 0 | - | ||||||||||
肌醇六乙酸酯 | v44 | 1254-38-2 | - | - | - | 2.57 | - | ||||||||||
甲基丙烯酸3-甲基戊酯 | v45 | 541-47-9 | - | 0.1 | - | 0 | - | ||||||||||
顺-9-十四碳烯酸异丁酯 | v46 | 16725-53-4 | - | 0 | - | 0.55 | - | ||||||||||
十二酸乙酯 | v47 | 18281-05-5 | - | 0.07 | 0.06 | 0 | - | ||||||||||
醛类 Aldehyde | 13-甲基十四醛 | v48 | 2485-71-4 | - | - | - | 1.09 | - | |||||||||
13-十八碳烯醛 | v49 | 58594-45-9 | - | - | - | 0.42 | - | ||||||||||
3-甲基-戊醛 | v50 | 15877-57-3 | - | - | - | 0 | 0.13 | ||||||||||
十六烷醛 | v51 | 629-80-1 | - | - | - | 0.57 | - | ||||||||||
十四醛 | v52 | 124-25-4 | - | - | - | 4.74 | - | ||||||||||
顺-13-十八烯醛 | v53 | 58594-45-9 | - | - | - | 0.71 | - | ||||||||||
正十五碳醛 | v54 | 2756-11-9 | 66.95 | 0.29 | 0.24 | 48.39 | 0.07 | ||||||||||
烷烃类 Alkanes | 十七烷 | v55 | 629-78-7 | - | - | - | 0.94 | - | |||||||||
2,6,10,15-四甲基-十七烷 | v56 | 18344-37-1 | - | - | - | 1.41 | - | ||||||||||
2-甲基-十九烷 | v57 | 1560-86-7 | - | - | - | 0.41 | - | ||||||||||
10,10-二甲基-2,6-二亚甲 二环[7.2.0]十一烷 | v58 | 87399-97-1 | - | 1.67 | 1.35 | - | 2.23 | ||||||||||
十四烷 | v59 | 629-59-4 | 0.44 | - | - | 1.62 | - | ||||||||||
十五烷 | v60 | 629-62-9 | 0.43 | - | - | 1.02 | - | ||||||||||
十六烷 | v61 | 544-76-3 | 0.37 | - | - | 2.15 | - | ||||||||||
酮类 Ketones | β-紫罗酮 | v62 | 79-77-6 | 3.81 | - | - | 3.02 | - | |||||||||
4-异丙基-5,10-二甲基 十氢萘-1,3-二酮 | v63 | 37830-90-3 | - | 0.1 | 0.09 | 0 | - | ||||||||||
5,6-环氧-β紫罗兰酮 | v64 | 20483-36-7 | - | - | - | 0.69 | - | ||||||||||
6,10,14-三甲基-2-十五烷酮 | v65 | 502-69-2 | - | - | - | 0.5 | - | ||||||||||
(Z)--香叶基丙酮 | v66 | 3796-70-1 | - | - | - | 0.65 | - | ||||||||||
酰胺类Amides | 己内酰胺 | v67 | 105-60-2 | - | - | - | 0.92 | - |
挥发物种类 Compound type | 双斑长跗萤叶甲取食前后玉米叶片挥发物名称 Name of the volatile substance released from maize leaves | |
---|---|---|
健康玉米叶片 Healthy Corn Leaf | 取食后的玉米叶片 Maize leaves after feeding | |
醇类 Alcohols | 1,3-丁二烯醇 | - |
7-十三烯醇(Z) | - | |
3-已基-1,5-辛二烯 | - | |
5-十三烯 | - | |
环庚醇 | - | |
- | 3,7,11-三甲基-1-十二烷醇 | |
- | (E)-2-十四碳烯-1-醇 | |
- | (E)-3-己烯-1-醇 | |
- | 3,7,7-三甲基双环[2.2.1]庚-2-甲基醇 | |
- | δ-杜松醇 | |
- | 橙花叔醇 | |
- | 异植物醇 | |
植醇 | 植醇 | |
β-红没药醇 | β-红没药醇 | |
萜类 Terpenes | 1,6-辛二烯 | - |
1-石竹烯 | - | |
2,3-二甲基-1,3-丁二烯 | - | |
3-已基1,5-辛二烯 | - | |
5-十三烯 | - | |
α-石竹烯 | - | |
反-Zα环氧化红没药烯 | - | |
异丁烯 | - | |
- | (1R)-(+)-α蒎烯 | |
- | (E)-β-法呢烯 | |
- | (Z,E)-α-法呢烯 | |
- | D-柠檬烯 | |
- | L-柠檬烯 | |
- | α-荜澄茄油烯 | |
- | α-蒎烯 | |
- | β-罗勒烯 | |
- | β-蒎烯 | |
- | β-檀香烯 | |
- | β月桂烯 | |
- | 倍半水芹烯 | |
- | 反式-α-香柠檬烯 | |
- | 葎草烯 | |
- | 顺-β-法呢烯 | |
- | 顺γ-红没药烯 | |
- | 新植二烯 | |
氧化石竹烯 | 氧化石竹烯 | |
绿叶化合物 Green leaf compounds | - | 3,5,5-三甲基-环己烯 |
正己醛 | - | |
- | 丁烯基环己烷 | |
- | 1,5,5-三甲基-6-亚甲基环己烯 | |
烷烃类 Alkanes | - | 2,6,10,15-四甲基-十七烷 |
- | 10,10-二甲基-2,6-二亚甲二环[7.2.0]十一烷 | |
2,6,6-三甲基3-(2-丙烯基)双环[3.1.1]庚烷 | - | |
十六烷 | 十六烷 | |
十四烷 | 十四烷 | |
十五烷 | 十五烷 | |
酮类 Ketones | - | (Z)-香叶基丙酮 |
- | 4-异丙基-5,10-二甲基十氢萘-1,3-二酮 | |
- | 5,6-环氧-β紫罗兰酮 | |
- | 6,10,14-三甲基-2-十五烷酮 | |
β-紫罗酮 | β-紫罗酮 | |
醛类 Aldehyde | 顺-11-十六醛 | - |
(Z)-9-肉豆蔻醛 | - | |
(Z)-9-十六碳烯醛 | - | |
1,3,4-三甲基-3环己烯-1-甲醛 | - | |
- | 13-甲基十四醛 | |
- | 13-十八碳烯醛 | |
- | 十六烷醛 | |
- | 十四醛 | |
- | 顺-13-十八烯醛 | |
正十五碳醛 | 正十五碳醛 | |
酯类 Esters | - | 2-甲基-壬基丙酸酯 |
- | 肌醇六乙酸酯 | |
邻苯二甲酸二异丁酯 | 邻苯二甲酸二异丁酯 | |
- | 十二酸乙酯 | |
- | 顺-9-十四碳烯酸异丁酯 | |
吡喃类Pyran | 2,5-二甲基四氢-2-甲醇吡喃 | - |
酸类Acids | 棕榈酸 | - |
表3 双斑长跗萤叶甲取食前后玉米叶片挥发物比较
Tab.3 Comparison of corn leaf volatiles before and after feeding by the M.signata
挥发物种类 Compound type | 双斑长跗萤叶甲取食前后玉米叶片挥发物名称 Name of the volatile substance released from maize leaves | |
---|---|---|
健康玉米叶片 Healthy Corn Leaf | 取食后的玉米叶片 Maize leaves after feeding | |
醇类 Alcohols | 1,3-丁二烯醇 | - |
7-十三烯醇(Z) | - | |
3-已基-1,5-辛二烯 | - | |
5-十三烯 | - | |
环庚醇 | - | |
- | 3,7,11-三甲基-1-十二烷醇 | |
- | (E)-2-十四碳烯-1-醇 | |
- | (E)-3-己烯-1-醇 | |
- | 3,7,7-三甲基双环[2.2.1]庚-2-甲基醇 | |
- | δ-杜松醇 | |
- | 橙花叔醇 | |
- | 异植物醇 | |
植醇 | 植醇 | |
β-红没药醇 | β-红没药醇 | |
萜类 Terpenes | 1,6-辛二烯 | - |
1-石竹烯 | - | |
2,3-二甲基-1,3-丁二烯 | - | |
3-已基1,5-辛二烯 | - | |
5-十三烯 | - | |
α-石竹烯 | - | |
反-Zα环氧化红没药烯 | - | |
异丁烯 | - | |
- | (1R)-(+)-α蒎烯 | |
- | (E)-β-法呢烯 | |
- | (Z,E)-α-法呢烯 | |
- | D-柠檬烯 | |
- | L-柠檬烯 | |
- | α-荜澄茄油烯 | |
- | α-蒎烯 | |
- | β-罗勒烯 | |
- | β-蒎烯 | |
- | β-檀香烯 | |
- | β月桂烯 | |
- | 倍半水芹烯 | |
- | 反式-α-香柠檬烯 | |
- | 葎草烯 | |
- | 顺-β-法呢烯 | |
- | 顺γ-红没药烯 | |
- | 新植二烯 | |
氧化石竹烯 | 氧化石竹烯 | |
绿叶化合物 Green leaf compounds | - | 3,5,5-三甲基-环己烯 |
正己醛 | - | |
- | 丁烯基环己烷 | |
- | 1,5,5-三甲基-6-亚甲基环己烯 | |
烷烃类 Alkanes | - | 2,6,10,15-四甲基-十七烷 |
- | 10,10-二甲基-2,6-二亚甲二环[7.2.0]十一烷 | |
2,6,6-三甲基3-(2-丙烯基)双环[3.1.1]庚烷 | - | |
十六烷 | 十六烷 | |
十四烷 | 十四烷 | |
十五烷 | 十五烷 | |
酮类 Ketones | - | (Z)-香叶基丙酮 |
- | 4-异丙基-5,10-二甲基十氢萘-1,3-二酮 | |
- | 5,6-环氧-β紫罗兰酮 | |
- | 6,10,14-三甲基-2-十五烷酮 | |
β-紫罗酮 | β-紫罗酮 | |
醛类 Aldehyde | 顺-11-十六醛 | - |
(Z)-9-肉豆蔻醛 | - | |
(Z)-9-十六碳烯醛 | - | |
1,3,4-三甲基-3环己烯-1-甲醛 | - | |
- | 13-甲基十四醛 | |
- | 13-十八碳烯醛 | |
- | 十六烷醛 | |
- | 十四醛 | |
- | 顺-13-十八烯醛 | |
正十五碳醛 | 正十五碳醛 | |
酯类 Esters | - | 2-甲基-壬基丙酸酯 |
- | 肌醇六乙酸酯 | |
邻苯二甲酸二异丁酯 | 邻苯二甲酸二异丁酯 | |
- | 十二酸乙酯 | |
- | 顺-9-十四碳烯酸异丁酯 | |
吡喃类Pyran | 2,5-二甲基四氢-2-甲醇吡喃 | - |
酸类Acids | 棕榈酸 | - |
[1] | 梁日霞, 王振营, 何康来, 等. 基于线粒体COⅡ基因序列的双斑长跗萤叶甲中国北方地理种群的遗传多样性研究[J]. 昆虫学报, 2011, 54(7):828-837. |
LIANG Rixia, WANG Zhenying, HE Kanglai, et al. Genetic diversity of geographic populations of Monolepta hieroglyphica(Motschulsky)(Coleoptera:Chrysomelidae)from North China estimated by mitochondrial COⅡ gene sequences[J]. Acta Entomologica Sinica, 2011, 54(7):828-837. | |
[2] | 张聪, 袁志华, 王振营, 等. 双斑长跗萤叶甲在玉米田的种群消长规律[J]. 应用昆虫学报, 2014, 51(3):668-675. |
ZHANG Cong, YUAN Zhihua, WANG Zhenying, et al. Population growth and decline patterns of the Monolepta hieroglyphica(Motschulsky)in maize fields[J]. Journal of Applied Entomology, 2014, 51(3):668-675. | |
[3] | 姜玉英, 曾娟. 警惕双斑长跗莹叶甲加重为害北方多种作物[J]. 中国植保导刊, 2008, 28(4):45-46. |
JIANG Yuying, ZENG Juan. Alert to the increase of Monolepta hieroglyphica(Motschulsky)infesting various crops in the north[J]. China Plant Protection, 2008, 28(4):45-46. | |
[4] | 王立仁, 刘斌侠, 付泓. 玉米田双斑长跗萤叶甲的发生为害情况与防治对策[J]. 陕西农业科学, 2006, 52(2):123-131. |
WANG Liren, LIU Binman, FU Hong. Occurrence and control measures of Monolepta hieroglyphica(Motschulsky)in maize fields[J]. Shaanxi Agricultural Sciences, 2006, 52(2):123-131. | |
[5] | 中国科学院动物研究所昆虫分类区系室叶甲组, 河北省张家口地区坝下农业科学研究所植保组. 双斑萤叶甲研究简报[J]. 昆虫学报, 1979, 22(1):115-117. |
Institute of Zoology, Chinese Academy of Sciences,Plant Protection Group, Zhangjiakou Regional Institute of Agricultural Science, Hebei Province. A brief study of the Monolepta hieroglyphica[J]. Acta Entomologica Sinica, 1979, 22(1):115-117. | |
[6] |
Ton J, D'Alessandro M, Jourdie V, et al. Priming by airborne signals boosts direct and indirect resistance in maize[J]. The Plant Journal for Cell and Molecular Biology, 2007, 49(1):16-26.
DOI URL |
[7] |
Paré P W, Tumlinson J H. Update on plant-insect interactions plant volatiles as a defense against insect herbivores[J]. Plant Physiology, 1999, 121(2):325-332.
PMID |
[8] |
Dicke M, van Loon J J A, Soler R. Chemical complexity of volatiles from plants induced by multiple attack[J]. Nature Chemical Biology, 2009, 5:317-324.
DOI PMID |
[9] | Aartsma Y. Herbivore-induced plant volatiles and tritrophic interactions from local to landscape scale[D]. Wageningen Vmiversity and Research, 2018. |
[10] | 严善春, 张丹丹, 迟德富. 植物挥发性物质对昆虫作用的研究进展[J]. 应用生态学报, 2003,(2):310-313. |
YAN Shanchun, ZHANG Dandan, CHI Defu. Research progress on the effects of plant volatiles on insects[J]. Chinese Journal of Applied Ecology, 2003,(2):310-313.
PMID |
|
[11] | 杜家纬. 植物-昆虫间的化学通讯及其行为控制[J]. 植物生理学报, 2001, 27(3):193-200. |
DU Jiawei. Chemical communication between plants and insects and its behavioral control[J]. Acta Photophysiologica Sinica, 2001, 27(3):193-200. | |
[12] | 蒋方一丁, 吕宝乾, 唐继洪, 等. 椰心叶甲危害对椰树心叶挥发物的影响[J]. 生物安全学报, 2020, 29(4):264-267. |
JIANGFANG Yiding, LYU Baoqian, TANG Jihong, et al. Variation of volatile constituents in fresh leaves of Coconut nucifera after the damage of Brontispa longissima Gestro[J]. Journal of Biosafety, 2020, 29(4):264-267 | |
[13] |
Gouinguené S, Alborn H, Turlings T C J. Induction of volatile emissions in maize by different larval instars of spodoptera littoralis[J]. Journal of Chemical Ecology, 2003, 29(1):145-162.
PMID |
[14] |
Frost C J, Mescher M C, Dervinis C, et al. Priming defense genes and metabolites in hybrid poplar by the green leaf volatile cis-3-hexenyl acetate[J]. The New Phytologist, 2008, 180(3):722-734.
DOI URL |
[15] |
Bate N J, Rothstein S J. C6-volatiles derived from the lipoxygenase pathway induce a subset of defense-related genes[J]. The Plant Journal, 1998, 16(5):561-569.
DOI URL |
[16] | 蔡晓明. 三种茶树害虫诱导茶树挥发物的释放规律[D]. 北京: 中国农业科学院, 2009. |
CAI Xiaoming. Release patterns of tea tree volatiles induced by three tea tree pests[D]. Beijing: Chinese Academy of Agricultural Sciences, 2009. | |
[17] |
Zhang S F, Wei J N, Guo X J, et al. Functional synchronization of biological rhythms in a tritrophic system[J]. PLoS One, 2010, 5(6):e11064.
DOI URL |
[18] |
张志虎, 陈静, 唐思琼, 等. 双斑长跗萤叶甲成虫对α-红没药醇等棉花和玉米挥发物的嗅觉行为反应[J]. 新疆农业科学, 2018, 55(6):1117-1123.
DOI |
ZHANG Zhihu, CHEN Jing, TANG Siqiong, et al. Olfactory Behavioral Response of the Monolepta hieroglyphica(Motschulsky)to volatiles of cotton and corn such as Dragosantol[J]. Xinjiang Agricultural Sciences, 2018, 55(6):1117-1123.
DOI |
|
[19] | 李鹏. 四种害虫对14种植物挥发物的EAG和行为反应[D]. 哈尔滨: 东北林业大学, 2012. |
LI Peng. EAG and behavioral responses of four pests to 14 plant volatiles[D]. Harbin:Northeast Forestry University, 2012. | |
[20] | 郭丹丹. 双斑长跗萤叶甲头部超微结构及其对寄主挥发物的响应[D]. 石河子: 石河子大学, 2018. |
GUO Dandan. Ultrastructure of the head of Monolepta hieroglyphica(motschulsky)and its response to host volatile compounds[D]. Shihezi: Shihezi University, 2018. | |
[21] |
Schilmiller A L, Howe G A. Systemic signaling in the wound response[J]. Current Opinion in Plant Biology, 2005, 8(4):369-377.
PMID |
[22] | 孙晓玲, 高宇, 陈宗懋. 虫害诱导植物挥发物(HIPVs)对植食性昆虫的行为调控[J]. 应用昆虫学报, 2012, 49(6):1413-1422. |
SUN Xiaoling, GAO Yu, CHEN Zongmao. Behavioral modulation of phytophagous insects by pest-induced plant volatiles(HIPVs)[J]. Chinese Journal of Applied Entomology, 2012, 49(6):1413-1422. | |
[23] | 娄永根, 程家安. 虫害诱导的植物挥发物:基本特性、生态学功能及释放机制[J]. 生态学报, 2000, 20(6):1097-1106. |
LOU Yonggen, CHENG Jiaan. Pest-induced plant volatiles:basic properties, ecological functions and release mechanisms[J]. Journal of Ecology, 2000, 20(6):1097-1106 | |
[24] | 刘永泉, 薛阿辉, 丁鹏, 等. 柑橘木虱胁迫对柠檬叶代谢组挥发性成分的影响[J]. 环境昆虫学报, 2019, 41(5):1089-1098. |
LIU Yongquan, XUE Ahui, DING Peng, et al. The effect of Diaphorina citri Kuwayama on metabonomics volatilecompounds in Citrus limon Burm leaves[J]. Journal of Environmental Entomology, 2019, 41(5):1089-1098. | |
[25] |
Turlings T C J, Erb M. Tritrophic interactions mediated by herbivore-induced plant volatiles:mechanisms, ecological relevance, and application potential[J]. Annual Review of Entomology, 2018, 63:433-452.
DOI PMID |
[26] | 叶盛, 王俊. 水稻虫害信息快速检测方法实验研究——基于电子鼻系统[J]. 农机化研究, 2010, 32(6):146-149,204. |
YE Sheng, WANG Jun. Experimental study on the rapid detection method of rice pest information based on electronic nose system[J]. Agricultural Mechanization Research, 2010, 32(6):146-149,204. | |
[27] |
金珊, 韩李伟, 叶乃兴, 等. 茶小绿叶蝉危害乌龙茶茶树品种的挥发物分析[J]. 热带作物学报, 2019, 40(3):576-582.
DOI |
JIN Shan, HAN Liwei, YE Naixing, et al. Analysis on volatiles of oolong tea varieties induced by empoasca sp[J]. Chinese Journal of Tropical Crops, 2019, 40(3):576-582.
DOI |
|
[28] | 郭井菲. 亚洲玉米螟取食玉米产生的诱导抗性机制研究[D]. 北京: 中国农业科学院, 2017. |
GUO Jingfei. Defenses mechanism induced by ostrinia furnacalis(Lepidoptera:Crambidae)feeding in maize[D]. Beijing: Chinese Academy of Agricultural Sciences, 2017. | |
[29] |
杨陈, 李伦, 阿尔曼, 等. 双斑长跗萤叶甲对13种挥发物的触角电生理反应[J]. 新疆农业科学, 2021, 58(7):1282-1290.
DOI |
YANG Chen, LI Lun, Aerman, et al. Electrophysiological responses of the monolept ahieroglyphica (Motschulsky)to 13 volatile compounds[J]. Xinjiang Agricultural Sciences, 2021, 58(7):1282-1290.
DOI |
|
[30] |
李伦, 杨陈, 陈静. 双斑长跗萤叶甲成虫粪便活性成分鉴定[J]. 新疆农业科学, 2020, 57(6):1103-1112.
DOI |
LI Lun, YANG Chen, CHEN Jing. Identification of active components in feces of monolepta hieroglyphica(motschulsky)[J]. Xinjiang Agricultural Sciences, 2020, 57(6):1103-1112.
DOI |
|
[31] |
Yan Z G, Wang C Z. Wound-induced green leaf volatiles cause the release of acetylated derivatives and a terpenoid in maize[J]. Phytochemistry, 2006, 67(1):34-42.
DOI URL |
[32] | 狄贵秋, 李杰, 卢旭弘, 等. 松瘿小卷蛾及其寄生蜂对落叶松挥发物的嗅觉行为反应[J]. 生态学报, 2019, 39(22):8675-8683. |
DI Guiqiu, LI Jie, LU Xuhong, et al. Olfactory responses of Cydia zebeana adults and their parasitic wasps to larch volatiles[J]. Acta Ecologica Sinica, 2019, 39(22):8675-8683. | |
[33] | 管维康, 王小云, 陆温, 等. 朱红毛斑蛾和花胸姬蜂对虫害诱导榕树挥发物的触角电位及行为反应[J]. 植物保护, 2022, 48(3):81-89. |
GUAN Weikang, WANG Xiaoyun, LU Wen, et al. Tactile potential and behavioural responses of the vermilion moth, Ficus vermiculata, and the flower-breasted gypsy wasp to pest-induced volatiles in Ficus[J]. Plant Protection, 2022, 48(3):81-89. | |
[34] | 周强, 徐涛, 张古忍, 等. 虫害诱导的水稻挥发物对褐飞虱的驱避作用[J]. 昆虫学报, 2003, 46(6):739-744. |
ZHOU Qiang, XU Tao, ZHANG Guren, et al. Repellent effects of herbivore-induced rice volatiles on Nilaparvata lugens Stal[J]. Acta Entomologica Sinica, 2003, 46(6):739-744. | |
[35] | 田厚军, 陈艺欣, 魏辉, 等. 芒果不同部位挥发物及横线尾夜蛾对其触角电位反应[J]. 生物安全学报, 2012, 21(2):142-147. |
TIAN Houjun, CHEN Yixin, WEI Hui, et al. Volatiles emitted by mango(Mangifera indica)plants and electrophysiological responses of Chlumetia transverse to such volatiles[J]. Journal of Biosafety, 2012, 21(2):142-147. | |
[36] |
Bruce T J A, Picktt J A. Perception of plant volatile blends by herbivorous insects-Finding the right mix[J]. Phytochemistry, 2011, 72(13):1605-1611.
DOI PMID |
[37] |
Geervliet J B F, Posthumus M A, Vet L E M, et al. Comparative analysis of headspace volatiles from different caterpillar-infested or uninfested food plants of pieris species[J]. Journal of Chemical Ecology, 1997, 23(12):2935-2954.
DOI URL |
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